Cloeodes redactus Waltz & McCafferty 1987
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|Cloeodes redactus Waltz & McCafferty 1987|
Cloeodes (Cloeodes) redactus Waltz & McCafferty 1987b: 204 (larva) (non C. redactus sensu Massariol et al. 2013 );? Cloeodes barituensis: Falcao, Salles & Hamada 2011 : Fig. 89 View FIGURES 86 – 91 (larva) (non C. barituensis Nieto & Richard 2008 ).
Material examined. PERU: Region Ucayali, Prov. Padre Abad, Aguaytia, Pampa Yurac , La Choza , 9°04'S, 75°31'W, 23.II.2006, coll. N. Kluge: 2 L/S ♀ GoogleMaps ; Region Loreto, Prov. Ucayali: 17 km NNE Contamana, Aguas Termales , 7°11'S, 74°57'W, 26.VII–4.VIII.2013, coll. N. Kluge & L. Sheyko: 5 L-S-I ♂, 1 L-S/I ♂, 7 L-S-I ♀, 8 larvae GoogleMaps ; ibid., Pampa Hermosa, Quebrada Huanganaico , 7°12'S, 75°22'W 19–20.VIII.2013, coll. N. Kluge & L. Sheyko: 3 L-S-I ♂, 7 larvae GoogleMaps .
Region Loreto, Prov. Maynas, Quebrada El Sabalo (right tributary of Rio Itaya) between Puente Itaya (57 km from Iquitos ) and San Joaquin (at Rio Amazon ), 4°16'S, 73°27'W, 1–16.II.2006, coll. N. Kluge: 1 L-S ♂, 1 L-S-I ♀, 2 L-S ♀, 7 larvae. GoogleMaps
Descriptions. Larva. CUTICULAR COLORATION ( Figs 58–62 View FIGURES 58 – 62 , 69–73 View FIGURES 69 – 75 ). Head with colorless and brown areas. Terga and pleura of thorax mainly brown with composite maculation, sterna colorless; in immature larva cuticle of fore protoptera colorless ( Figs 58–59 View FIGURES 58 – 62 ), before molt cuticle of protoptera becomes brown ( Fig 72 View FIGURES 69 – 75 ). Legs light with or without diffusive darker macula at middle of femur, knee articulation and base of tarsus ( Figs 60–62 View FIGURES 58 – 62 ). Most of abdominal terga brown with lighter areas; pigmentation variable, but pigmentation of tergum II always less extensive than pigmentation of tergum III (Figs 5 8–59, 69–71); terga IV and VIII usually lighter than others, often at most part non-pigmented ( Fig. 70 View FIGURES 69 – 75 ), sometimes entirely non-pigmented ( Fig. 71 View FIGURES 69 – 75 ), rarely as dark as others ( Fig. View FIGURES 69 – 75
69); tergum IX more or less pigmented; tergum X well-pigmented. Abdominal sterna light, sometimes with brown lateral markings; paraprocts brown, as remainder segment X. Caudalii lighter, with brown band near apex ( Fig. 73 View FIGURES 69 – 75 ).
HYPODERMAL COLORATION. Black transverse stripes on posterior margins of abdominal terga I–III and V–VII often present ( Figs 58–59 View FIGURES 58 – 62 ), sometimes absent (as in imago). Female larva just before molt to subimago with lateral triangular brown maculae on abdominal sterna (as in imago—Fig. 86).
SHAPE AND SETATION. Frontal suture pointed, rectangular; face relatively long; in female larva eyes brought together (as in Fig. 6 View FIGURES 6 – 12 ). Labrum equally wide at base and at middle or little wider at base, with pair of submedian long setae and 2 pairs of sublateral long setae (as in Fig. 12 View FIGURES 6 – 12 ; Roback 1966: Fig. 31 View FIGURES 26 – 31 ). Prostheca of left mandible with 3 blunt processes and usually 4 (rarely 3 or 5) pointed processes; median margin of left mandible proximad of prostheca straight or slightly convex, smooth ( Figs 63, 65 View FIGURES 63 – 68 ). Prostheca of right mandible directed medially-proximally, bifurcate, with longest branch directed proximally; median margin of right mandible proximad of prostheca straight or slightly convex, smooth or with serration, without seta-like processes ( Figs 64, 66 View FIGURES 63 – 68 ). Stipes of maxilla proximally with 3–4 setae on ventro-medial side. Maxillary canines and distal dentiseta stout; distal dentiseta widened, with apex somewhat hooked toward canines (as in Fig. 8 View FIGURES 6 – 12 ; Roback 1966: Fig. 32 View FIGURES 32 – 36 ). Maxillary palp about as long as maxilla, 2-segmented ( Roback 1966: Fig. 34 View FIGURES 32 – 36 ). Labium with glossae and paraglossae subequal, both narrowed apically (as in Fig. 14 View FIGURES 13 – 17 ). Glossa with 6–8 setae in dorso-lateral row; ventrally with irregularly arranged setae in proximal part and 6–9 setae forming ventro-median row. Paraglossa with lateroapical setae forming one regular row and few (2–4) setae just dorsad of it; with 6–7 setae in ventro-median row; with 3–4 setae in dorso-median row. Distal segment of labial palp not widened apically, with median margin subequal to lateral margin (as in Figs 13, 14 View FIGURES 13 – 17 ; Roback 1966: Fig. 36 View FIGURES 32 – 36 ).
All thoracic terga without protuberances. Metanotum without vestiges of hind protoptera ( Fig. 61 View FIGURES 58 – 62 ). Fore legs largest, middle legs smaller and hind legs smallest; on fore leg tarsus much longer than tibia, on middle and hind legs tarsus subequal to tibia; claws not elongated, much less than half of tarsal length ( Table 4; Figs 60–62 View FIGURES 58 – 62 ). Femur parallel-sided, with outer margin straight and inner margin slightly convex; apical projection equally short on femora of all leg pairs (as in Fig. 16 View FIGURES 13 – 17 ). Outer side of femur with row of 3–6 small spatulate setae; apical projection with 2 such setae (as in Fig. 16 View FIGURES 13 – 17 ). Inner and dorsal side of femur with few stout setae, which are several times smaller than setae on dorsal side. Patella-tibial suture and proximal arc of long setae with contiguous sockets equally developed on tibiae of all leg pairs; inner margin of tibia with irregular small stout pointed setae, apically with 2 larger setae (as in Fig. 16 View FIGURES 13 – 17 ). Outer-apical seta blunt, reaching apex of tibia ( Fig. 68 View FIGURES 63 – 68 ). Dorsal side of each tarsus with long fine setae forming 2 longitudinal rows ( Fig. 68 View FIGURES 63 – 68 ). Claw shorter than 1/2 of tarsus length ( Fig. 68 View FIGURES 63 – 68 ).
Scales on abdominal terga and sterna numerous, short, semicircular, colorless and delicate. Posterior margin of abdominal tergum I often entirely smooth, rarely with few irregular short denticles; posterior margins of all terga II–X often with regular long pointed denticles, rarely tergum II without denticles or with few irregular short denticles; on tergum IX, portion of posterior margin behind pair of submedian setae, lacks denticles and projected backward (as in Fig. 18 View FIGURES 18 – 25 ). Posterior margins of abdominal sterna I–IV smooth; posterior margins of sterna V–VIII with regular pointed denticles, shorter and lighter than denticles on terga. Each sternum II–VI with pair of regular transverse arched rows of long, fine, simple (not furcate) setae with closely contiguous sockets (as in Fig. 20 View FIGURES 18 – 25 ). Tergalius I widest at distal half, with costal and anal margins convex; tergalius II widest near middle, with anal margin more convex than costal margin; tergalii III–IV widest at proximal half, with anal margin more convex than costal margin ( Figs 44–57 View FIGURES 37 – 57 ). All tergalii with ribs only on proximal parts of costal and anal margins, without middle rib. Margins bearing costal and anal ribs smooth (not serrated); margin free of ribs slightly notched, with small seta in each notch. Paraprocts with anterior median apodeme small, soft and colorless; each paraproct with 7–8 long pointed denticles on posterior margin, with dense scales as on sterna and terga ( Fig. 67 View FIGURES 63 – 68 ). Each swimming seta of caudalii sharply divided into dark proximal part and colorless delicate distal part ( Fig. 73 View FIGURES 69 – 75 ). Ventral and dorsal side of paracercus and lateral sides of cerci in middle part with one or several long pointed denticles on each 4th segment.
DEVELOPING MALE GENITALS. In last larval instar, developing subimaginal gonostyli folded under larval cuticle in « Nigrobaetis -type» pose, with 2nd segment bent by convexity medially, and 3rd segment directed caudally ( Figs 86, 89 View FIGURES 86 – 91 ).
RESPIRATORY MOVEMENTS. Larva does not make respiratory vibration by the body (unlike C. vibratorius ). Tergalii unable to respiratory movements.
Subimago. CUTICULAR COLORATION. Cuticle with light brown and colorless areas. Head colorless, only antennae slightly tinged with pale brownish. Pronotum with light brown and colorless areas. Mesonotum at most brown, medioparapsidal suture contrastingly colorless, other sutures darkened ( Fig. 74 View FIGURES 69 – 75 ). Thoracic pleura with light brown and colorless areas; lateropostnotal crest narrowly colored by dark brown, diffusively bordered by light brown ( Fig. 75 View FIGURES 69 – 75 ). Base of wing light brown, other wing cuticle lighter. Legs at most colorless, tarsi slightly tinged with pale brownish; distal margin of each tarsal segment narrowly bordered by brown. Abdominal terga slightly colored with light brownish, microtrichiae brown. Abdominal sterna lighter. Cerci colorless.
HYPODERMAL COLORATION. As in imago.
TEXTURE. On all legs of both sexes last tarsal segment entirely covered by pointed microlepides; other tarsal segments at most part covered by blunt microlepides, apically by pointed microlepides ( Table 1).
Tarsal spines of subimago and imago. Tarsus of fore leg in male and female without apical spines. Tarsus of middle and hind leg of both sexes with 1 apical spine on initial 3rd tarsomere (next after 1st+2nd tarsomere) ( Table 1).
Imago, male. COLORATION ( Fig. 76–81 View FIGURES 76 – 81 ). Head ocher. Turbinate eyes orange or yellow. Thorax ocher with light brown and white markings. Wing membrane colorless, veins pale ocher or colorless, basal part of costal and subcostal veins and part of costal brace brown. Femora of all legs ocher, with brown or reddish subapical band (hypodermal coloration); main trachea can be colored with dark brown; fore femur also with brown macula apically. Tibiae of all legs pale ocher, with or without hypodermal brownish coloration on inner side. Tarsi of all legs pale ocher, on fore leg cuticle of first segment can be tinged with brownish. Claws as light as tarsus. Abdominal segments I–III and V–VI colorless; segment IV either also colorless ( Fig. 77 View FIGURES 76 – 81 ), or with tergum tinged with pale yellowish ( Figs 76, 78 View FIGURES 76 – 81 ); segments VII–X pale yellowish; terga always without median markings; terga I– VII with contrasting dark paired transverse stripes near posterior margin: stripes on tergum I largest, stripes on terga IV and VII smallest. Color of these stripes varies from black to brown to light reddish; size of stripes varies from wide and connected medially to small and dot-shaped; stripes can be completely absent on terga IV and/or VII, or on terga II–IV and/or VII, or present on tergum I only. Lateral tracheal trunks and bases of sternal tracheae contrastingly colored by black or brown. Cerci entirely colorless, whitish.
STRUCTURE. Turbinate eyes with facetted surface wider than stem ( Figs 76–79 View FIGURES 76 – 81 ). Pterostigma with 3–5 complete and incomplete oblique veins. Two marginal intercalaries nearly in each space (except for few posterior ones), length of most intercalaries less than distance between adjacent longitudinal veins (as in Fig. 32 View FIGURES 32 – 36 ). Hind wing absent. Middle and hind tarsus long (about 0.7–0.8 of tibia length), its 1st+2nd segment subequal to 3rd, 4th and 5th segments together ( Table 4). Genitals as in Figs 88, 91 View FIGURES 86 – 91 : unistyligers cylindrical; median sterno-styligeral muscle slender, consists of separate bunches, which are either convergent V-shaped ( Fig. 88 View FIGURES 86 – 91 ), or crossed ( Fig. 91 View FIGURES 86 – 91 ).
Imago, female. COLORATION ( Figs 82–85 View FIGURES 82 – 85 ). Coloration of head, thorax, wings and legs similar to that of male. Abdomen ocher, terga VI and VII darker; terga with contrasting dark paired transverse stripes as in male; besides this, lateral parts of terga, pleura and lateral parts of sterna with composite brown and/or reddish maculation; particularly, at least each sternum II–VII with pair of contrasting brown lateral triangular maculae widened toward anterior margin of sternum.
STRUCTURE. Eyes relatively large, elevated above head surface ( Fig. 83 View FIGURES 82 – 85 ). Wing either with one marginal intercalary in some spaces and no one marginal intercalary in other spaces, or without marginal intercalaries at all; in other respects wing venation as in male ( Figs 82, 85 View FIGURES 82 – 85 ).
Egg. Oval, 0.1 mm length. Chorion irregularly crumpled, without relief ( Fig. 146 View FIGURES 145 – 146 ).
Dimension. Fore wing length: male and female 4 mm.
Variability. Among specimens examined, two forms can be distinguished.
The first form («pallidus») includes specimens collected near Puerto Prado (6 male and 4 female imagoes and subimagoes associated with larvae). They have transverse stripes on abdominal terga at most light brown or reddish, poorly expressed or non-expressed ( Fig. 79 View FIGURES 76 – 81 ); larvae of all these 10 specimens have prostheca of left mandible with 3 blunt processes and 3 pointed processes ( Fig. 65 View FIGURES 63 – 68 ).
The second form («nigrostriatus») includes specimens collected near Contamana, Pampa Hermosa and Puente Itaya (11 male and 11 female imagoes and subimagoes associated with larvae). They have transverse stripes on imaginal abdominal terga at most well expressed, contrastingly dark brown or black ( Figs 76–78 View FIGURES 76 – 81 ); among them, 18 larvae have prostheca of left mandible with 3 blunt processes and 4 pointed processes ( Fig. 63 View FIGURES 63 – 68 ); 2 specimens have prostheca of left mandible with 3 blunt processes and 5 pointed processes; 2 specimens (male and female imagoes reared from larvae near Contamana) have prostheca of left mandible with 3 blunt processes and 3 pointed processes.
Among 2 specimens from Aguaytia (mature female larvae with developed subimaginal parts) one has features of the form «pallidus» (small brown hypodermal stripes on abdominal terga and left prostheca with 3+3 processes) and another has features of the from «nigrostriatus» (long black hypodermal stripes on abdominal terga and left prostheca with 3+4 processes).
The form «pallidus» well agrees with the original description of C. redactus based on larvae from Tingo Maria, for which dark hypodermal stripers on larval abdominal terga are not reported, and left prostheca is figured with 3 blunt processes and 3 pointed processes ( Roback 1966: Fig. 33 View FIGURES 32 – 36 ).
I could not find other differences between these two forms, and regard them to be conspecific.
Comparison. Female imago, female subimago and female mature larva of C. redactus can be recognized by presence of contrasting brown hypodermal triangular maculae on abdominal sterna. Imago and subimago of both sexes can be recognized by having contrasting dark transverse lines or paired stripes on posterior margins of abdominal terga ( Figs 76–78 View FIGURES 76 – 81 , 84 View FIGURES 82 – 85 ), but in some individuals these markings are lighter ( Fig. 79 View FIGURES 76 – 81 ) or absent. Larva can be distinguished from other species by the characters given in the key (see above).
Discussion. Preliminary description (without formal species name) was based on 6 larvae from three rivers near Tingo Maria in Peru ( Roback 1966). Formal original description was based on 2 of these larvae from the river Cantaria at Puente Perez ( Waltz & McCafferty 1987b).
Larvae described above, well agree with the original description; they were collected in various localities in Peru, and two specimens were collected near Aguaytia, about 50 km from the type locality.
McCafferty & Lugo-Ortiz (1996: 19–28) reported “ C. redactus ” from Honduras without more details.
Nieto and co-authors ( Nieto & Richard 2008; Nieto & Emmerich 2011) keyed larva of “ C. redactus ” as having denticles on posterior margin on abdominal tergum I, but did not report origin of this information. Actually, denticles on abdominal tergum I are absent or vestigial.
Massariol et al. (2013) described larvae and female imago determined by them as “ C. redactus ”. Description of this female imago is based on one female imago reared from larva in Reserva Ducke in Manaus, while description of larvae are based on 20 larvae collected in Igarape do Anta in Barcelos (F.F. Salles, personal communication). Photos of these larvae contradict the original description of C. redactus : in C. redactus “abdomen dark brown with light pattern ... tergites 4 and 8 lighter” ( Roback 1966) ( Waltz & McCafferty 1987b: Fig. 43 View FIGURES 37 – 57 ), while on the photos abdominal terga I–IV and VII–VIII nearly lack cuticular pigmentation, and only terga V–VI and IX–X have brown cuticular pigmentation ( Massariol et al. 2013: Figs 5, 6). Female imago ascribed by Massariol et al. (2013) to C. redactus has abdominal terga with red and orange median markings and sterna washed with orange with lighter sigilla ( Massariol et al. 2013: Fig. 31–32 View FIGURES 26 – 31 View FIGURES 32 – 36 ), while female imagoes reared by me, have no such red and orange coloration, but have contrasting brown lateral spots on abdominal sterna ( Figs 82–86 View FIGURES 82 – 85 View FIGURES 86 – 91 ). Thus, larvae and female imago described by Massariol et al. (2013) under the name “ Cloeodes redactus ”, do not belong to this species, and possibly represent a new species.
Falcao et al. (2011) reported larvae of C. barituensis Nieto & Richard 2008 from Roraima, Brazil and gave photos of total dorsal view and labium ( Falcao et al. 2011: Figs 89, 90 View FIGURES 86 – 91 ). Coloration of this larva markedly differs from described for C. barituensis . Judging by cuticular coloration (mainly brown with lighter blanks and lightest abdominal segments IV and VIII) and hypodermal coloration (dark transverse stripes on posterior margins of abdominal terga) larva from Roraima can belong to C. redactus .
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